Observed variation in species richness on ecological gradients and between regions has attracted several different explanations. Climatic factors, such as energy availability, precipitation and mean annual temperature, are frequently cited to explain differences in species richness (Wright et al. 1993). The relative amount of variation in these variables may be of importance as well as absolute values (Stevens 1989, 1992). Alternatively, the history of colonization and extirpation have been used to explain regional variation in diversity (Guo et al. 1998; Latham & Ricklefs 1993a, b). Area is an important factor, for example on an elevational gradient the tops of mountains are not only cooler than the bottoms, but they are also much smaller (Rahbek 1997).